• Ant association originated three times independently in blues and metalmarks

Summary

Butterflies (Papilionoidea), with over 18,000 described species, have captivated naturalists and scientists for centuries. They play a central role in the study of speciation, community ecology, biogeography, climate change, and plant-insect interactions and include many model organisms and pest species. However, a robust higher-level phylogenetic framework is lacking. To fill this gap, we inferred a dated phylogeny by analyzing the first phylogenomic dataset, including 352 loci (> 150,000 bp) from 207 species representing 98% of tribes, a 35-fold increase in gene sampling and 3-fold increase in taxon sampling over previous studies. Most data were generated with a new anchored hybrid enrichment (AHE) gene kit (BUTTERFLY1.0) that includes both new and frequently used (e.g., [6]) informative loci, enabling direct comparison and future dataset merging with previous studies. Butterflies originated around 119 million years ago (mya) in the late Cretaceous, but most extant lineages diverged after the Cretaceous-Paleogene (K-Pg) mass-extinction 65 mya. Our analyses support swallowtails (Papilionidae) as sister to all other butterflies, followed by skippers (Hesperiidae) + the nocturnal butterflies (Hedylidae) as sister to the remainder, indicating a secondary reversal from diurnality to nocturnality. The whites (Pieridae) were strongly supported as sister to brush-footed butterflies (Nymphalidae) and blues + metalmarks (Lycaenidae and Riodinidae). Ant association independently evolved once in Lycaenidae and twice in Riodinidae. This study overturns prior notions of the taxon’s evolutionary history, as many long-recognized subfamilies and tribes are para- or polyphyletic. It also provides a much-needed backbone for a revised classification of butterflies and for future comparative studies including genome evolution and ecology.

Conclusion:

Our study confirms the power of phylogenomic approaches to resolve challenging arthropod phylogenetic relationships. Adding more than 340 genes to the 10 used previously and tripling the number of taxa included in previous studies confirmed some formerly poorly supported nodes and indicated many novel relationships. A well-supported phylogeny with broad coverage across tribes enables tests of existing hypotheses about higher-level relationships and identification of areas needing further study. Critically, it also serves as a needed scaffold for testing entirely new questions about the tempo and mode of butterfly evolution, such as associations between butterfly and plant clades and the impact of the K-Pg mass-extinction event. Moreover, the phylogeny provides the needed framework for broad comparative studies of the origins of key innovations, such as caterpillar-ant symbioses and other hypothesized drivers of lineage diversification, that have shaped the evolution of this highly studied insect group.

Four heteropod lizard trackways discovered in the Hasandong Formation (Aptian-early Albian), South Korea assigned toSauripeshadongensis, n. ichnogen., n. ichnosp., which represents the oldest lizard tracks in the world. Most tracks are pes tracks (N = 25) that are very small, average 22.29 mm long and 12.46 mm wide. The pes tracks show “typical” lizard morphology as having curved digit imprints that progressively increase in length from digits I to IV, a smaller digit V that is separated from the other digits by a large interdigital angle. The manus track is 19.18 mm long and 19.23 mm wide, and shows a different morphology from the pes. The predominant pes tracks, the long stride length of pes, narrow trackway width, digitigrade manus and pes prints, and anteriorly oriented long axis of the fourth pedal digit indicate that these trackways were made by lizards running bipedally, suggesting that bipedality was possible early in lizard evolution.

Figure 1 Photograph and drawing of lizard trackways on the block.

Figure 5 A reconstruction of a lizard running bipedally chased by the pterosaur Pteraichnus koreanensis, based on the trackway (Drawn by Chuang Zhao).

Systematic ichnology

Order Squamata Oppel, 1811

Sauripeshadongensis ichnogen. et ichnosp. nov.

Etymology: Ichnogenus named from ancient Greek “sauros” (lizard) and “pes” (foot). Ichnospecies named after Hadong County that yielded the holotype.

Type locality and horizon: Hasandong Formation, Lower Cretaceous (Aptian-early Albian), an abandoned quarry next to Hadong power plant, Hadong County, South Gyeongsang Province, South Korea.

Diagnosis: Quadrupedal tracks; manus prints are medial to the pes prints; the pes prints are larger than the manus prints; plantigrade and pentadactyl pes prints are longer than wide; the digit length progressively increasing from digits I to IV (ectaxonic); digit V is oriented more laterally and offset from other digits; digit imprint IV is more than twice the length of the metatarsal impression; plantigrade and pentadactyl manus print has similar length and width dimensions; digits II and IV are shorter than digit III (mesaxonic); the interdigital angle between digits I and V of the manus is larger than that of the pes.

A herd of Saurolophus angustirostris moves along a river bank after a storm in the Cretaceous Nemegt Basin. The feet of the large herbivores sink into the soft sediment crushing the skull of a Tarbosaurus bataar that was lying in the mud.

Illustration based on specimen MPC-D107/05 collected at the Nemegt locality (Nemegt Formation) and discovered by J.Ed. Horton. Artwork by Davide Bonadonna.

Fig. 1: A herd of Saurolophus angustirostris moves along a river bank after a storm in the Cretaceous Nemegt Basin. The feet of the large herbivores sink into the soft sediment crushing the skull of a Tarbosaurus bataar that was lying in the mud. Illustration based on specimen MPC-D107/05 collected at the Nemegt locality (Nemegt Formation) and discovered by J.Ed. Horton.

We describe a new species of Liolaemus of the L. alticolor-bibronii group of the subgenus Liolaemus sensu stricto. We studied meristic, morphometric and qualitative pattern characters. Statistical tests were performed in order to evaluate morphological differences among the candidate species and the most closely geographically distributed species. Molecular analyses of Cyt-b mitochondrial gene were performed in order to estimate the position of the new species in relation to other taxa. We also recorded natural history data such as habitat, behavior, reproductive state, diet, and body temperature. Liolaemus absconditus sp. nov. differs from other species of Liolaemus in presenting a distinct combination of morphological character states of lepidosis and color pattern, being phylogenetically close to Liolaemus tandiliensis, Liolaemus gracilis and Liolaemus saxatilis. The new species is a saxicolous and endemic lizard of the Tandilia Mountain Range System of Buenos Aires Province.

A new species of African reed frog (genus Hyperolius Rapp, 1842) is described from the Coastal Forests of the Eastern Africa Biodiversity Hotspot in northeastern Mozambique. It is currently only known from less than ten localities associated with the Mozambican coastal pans system, but may also occur in the southeastern corner of Tanzania. Phylogenetic reconstructions using the mitochondrial 16S marker revealed that it is the sister taxon of Hyperoliusmitchelli (>5.6% 16S mtDNA sequence divergence) and forms part of a larger H. mitchelli complex with H. mitchelli and H. rubrovermiculatus. The new species is distinguished from other closely related Hyperolius species by genetic divergence, morphology, vocalisation, and dorsal colouration.

Euryhaline Cichlid fish of the species Coptodon guineensis are present in different water holes situated in a dried depression in the desert in the extreme South of Morocco, the Sebkha of Imlili. A genetic survey of this population, using complete sequences of the ND2 gene (mtDNA) and sixteen microsatellite loci, revealed that the fish in the sebkha did not form a single population, but rather a metapopulation. This metapopulational structure may be regarded as good news from the point of view of the conservation of fish in the sebkha. Although small individual populations may have short, finite life spans, the metapopulation as a whole is more stable, because immigrants from one population are likely to re-colonize the habitat, left open by the extinction of another.

Keywords: Tilapia; Conservation; Endangered species

Fig. 2. Views of the sebkha and the fish present in the water holes.
A. View from the top of the sand dunes. B. Band of plants surrounding the sebkha. C. Sandy soil with salt crystallization. D. Permanent water holes (two are clearly visible, five are indicated with arrows). E. Fish in a permanent hole. F. Close up of two specimens of C. guineensis from the sebkha (male above, female below).

Hoya phuluangensis Kidyoo, a new species from northeastern Thailand is here described and compared to the similar species, H. rostellata and H. siamica. All three species have glabrous stems and branches, glabrous coriaceous leaves, adaxially puberulent ovate corolla lobes with an acute apex, and flat to slightly erect coronal scales with an obtuse or rounded apex. However, H. phuluangensis differs from the other two species in the following characters: flowers with a shallow cup-shaped corolla tube and a corona diameter measuring less than half of the corolla tube diameter. Full description of H. phuluangensis is provided, together with line drawings and photographs. In addition, three new combinations and two new names in the genus Vincetoxicum, namely V. indicum (Burm.f.) Mabb. var. glabrum (Decne.) A. Kidyoo, V. kerrii (Craib) A. Kidyoo, V. sootepense (Craib) A. Kidyoo, V. lindleyi A. Kidyoo and V. potamophilum A. Kidyoo, are proposed.

Aroonrat Kidyoo and Manit Kidyoo. 2018. A New Species of Hoya (Marsdenieae), Three New Combinations and Two New Names in Vincetoxicum (Asclepiadeae) from Thailand. Taiwania. 63(1); 25-31. DOI: 10.6165/tai.2018.63.25

A new genus of flatid planthoppers (Hemiptera: Fulgoromorpha: Flatidae) is described from the island of Socotra (Yemen): Socoflata gen. nov., for Socoflata aurolineata sp. nov. and Socoflata histrionica sp. nov. (type species). Habitus, male and female external and internal genital structures of the new species are illustrated and diagnosed. Both Socoflata species are abundant and syntopic in the evergreen montane woodland and dwarf shrubland at high elevations in the Hagher mountains in central Socotra and are likely endemics of this area.

A new species of freshwater crab of the family Trichodactylidae, genus Rodriguezia Bott, 1969 is described from Grutas de Agua Blanca in southern Tabasco, Mexico. Rodriguezia is a genus endemic to northern Chiapas and southern Tabasco, distributed over a small area of 70 km.Rodriguezia adani n. sp., the third species of the genus, occurs north of its two congeners, being stygobitic with obvious adaptations to cave life. It can be distinguished from R. villalobosi, an epigean species, by the absence of eyes, lack of pigmentation and elongation of the pereiopods; and from R. mensabak by having less elongated pereiopods relative to carapace breadth, an extremely reduced ocular peduncle, and a smaller adult size.

Trachemys (Testudines: Emydidae) represents one of the most well-known turtle genera today. The evolution of Trachemys, while being heavily documented with fossil representatives, is not well understood. Numerous fossils from the late Hemphillian Gray Fossil Site (GFS) in northeastern Tennessee help to elucidate its evolution. The fossil Trachemys at the GFS represent a new species. The new taxon, Trachemys haugrudi, is described, and currently represents the most thoroughly described fossil emydid species known. A phylogenetic analysis, including 31 species, focusing on the subfamily Deirochelyinae is performed that includes the new fossil species, along with numerous other modern and fossil deirochelyine species, representing the first phylogenetic analysis published that includes several fossil deirochelyines. The phylogenetic analysis, utilizing morphological evidence, provides monophyletic clades of all modern deirochelyines, including Chrysemys, Deirochelys, Pseudemys, Malaclemys, Graptemys, and Trachemys. A strict consensus tree finds the recently described fossil species Graptemys kerneri to be part of a clade of Graptemys + Malaclemys. Three fossil taxa, including one previously referred to Pseudemys (Pseudemys caelata) and two to Deirochelys (Deirochelys carri and Deirochelys floridana) are found to form a clade with modern Deirochelys reticularia reticularia, with D. floridana sister to the other members of the clade. Chrysemys is found to be part of a basal polytomy with Deirochelys in relation to other deirochelyine taxa. Two fossil taxa previously referred to Chrysemys (Chrysemys timida and Chrysemys williamsi) form a paraphyly with the modern Chrysemys picta picta and Deirochelys, and may be referable to distinct genera. Additionally, fossil taxa previously attributed to Trachemys (Trachemys hillii, Trachemys idahoensis, Trachemys inflata, and Trachemys platymarginata) and T. haugrudi are found to form a clade separate from clades of northern and southern Trachemys species, potentially suggesting a distinct lineage of Trachemys with no modern survivors. Hypotheses of phylogenetic relationships mostly agree between the present study and previous ones, although the inclusion of fossil taxa provides further clues to the evolution of parts of the Deirochelyinae. The inclusion of more fossil taxa and characters may help resolve the placement of some taxa, and further elucidate the evolution of these New World turtles.

Figure 7: Trachemys haugrudi, paratype skull (ETMNH–3562) in dorsal view. (A) Dorsal portion in dorsal view; (B) ventral portion in dorsal view; (C) reconstruction of skull in dorsal view. Area shaded gray is not preserved and has been reconstructed. Dotted lines represent sutures that were not clear in the specimen. Skull has been reconstructed in the slightly deformed state the specimen is in in real life.

Etymology: The specific name honors Shawn Haugrud, preparator at the GFS who spent countless hours working on many of the specimens cited within and who helped piece this ancient turtle back together.

Diagnosis: Trachemys haugrudi is placed in the Emydidae due to the absence of musk ducts, inframarginals reduced to two, normal hexagonal neurals 2–8 (also occurs in a few batagurids (=geoemydids); e.g., Mauremys), and costal-inguinal buttress confined to C5. It is placed in the Deirochelyinae due to distinct lack of pectoral overlap of the entoplastron and lack of a hingable plastral lobe with ligamentous bridge connection (also present in some emydines). Diagnosed as a member of the genus Trachemys by features discussed by Seidel & Jackson (1990), including the combination of: a posteriorly strongly serrated oval carapace; a vertebral keel; low longitudinal ridges (mainly on pleurals (and costals)); alternating seams of the vertebral and pleural scutes that do not align; ......

Figure 13: Life reconstruction of Trachemys haugrudi during the late Hemphillian at the Gray Fossil Site in eastern Tennessee.
Several taxa that would have lived alongside T. haugrudi are also shown, including Caudataindeterminate, Tapirus polkensis, cf. Machairodus sp., and Pristinailurus bristoli (Wallace & Wang, 2004; Boardman & Schubert, 2011b; Schubert, 2011; Jasinski, 2013a). Artwork by Mary P. Williams, with permission.

Steven E. Jasinski. 2018. A New Slider Turtle (Testudines: Emydidae: Deirochelyinae: Trachemys) from the late Hemphillian (late Miocene/early Pliocene) of eastern Tennessee and the Evolution of the Deirochelyines. PeerJ. 6:e4338. DOI: 10.7717/peerj.4338

Satanoperca curupira, new species, is described from the rio Madeira basin in the State of Rondônia, Brazil. It is distinguished from all congeners by the following combination of characters: 3–7 dark-brown oblique stripes on the lachrymal (vs. 2 well-defined dark-brown stripes, or dark-brown stripes absent) and an irregular pattern of dark-brown stripes on cheek and opercular series (vs. cheek without dark-brown markings or with light-beige rounded spots). According to meristic and color pattern characters, the new species is considered a member of the S. jurupari species group, and is syntopic with S. jurupari, which is widespread in the Amazon basin. The restricted geographical range of the new species is congruent to that observed for some other Satanoperca species.

Distribution. Satanoperca curupira is known from the rio Madeira basin in Brazil, occurring in the main channel of the rio Madeira and in several tributaries draining the Brazilian shield (rio Jaciparaná, rio Jamari, rio Jaru and rio Machado), and also at the rio Roosevelt (a tributary of the rio Aripuanã), and at the rio São Luis (a tributary of the rio Guaporé) (Fig. 5).

Etymology. The specific name curupira refers to a mythological creature of Brazilian folklore that protects the forest and its inhabitants, punishing those who hunt for pleasure or who kill breeding females or defenseless juveniles (Pereira 1994). The Curupira legend reveals the relationship between the indigenous people and the forest: it is not about exploration and indiscriminate use, but respect for life. A noun in apposition.

Banded Knifefishes (Gymnotus, Gymnotidae) comprise the most species-rich, ecologically tolerant (eurytopic), and geographically widespread genus of Neotropical electric fishes (Gymnotiformes), with 40 valid species occupying most habitats and regions throughout the humid Neotropics. Despite substantial alpha-taxonomic work in recent years, parts of the genus remain characterized by taxonomic confusion. Here we describe and delimit species of the G. carapo and G. tigre clades from the southern Neotropics, using body proportions (caliper-based morphometrics), fin-ray, scale and laterosensory-pore counts (meristics), quantitative shape differences (geometric morphometrics), osteology, color patterns and electric organ discharges. We report these data from 174 Gymnotus specimens collected from 100 localities throughout the southern Neotropics, and delimit species boundaries in a multivariate statistical framework. We find six species of the G. carapo clade (G. carapo australis, Gymnotus cuia n. sp., G. chimarrao, G. omarorum, G. pantanal, and G. sylvius), and two species of the G. tigre clade (G. inaequilabiatus and G. paraguensis) in the southern Neotropics. The new species G. cuia is readily distinguished from the morphologically similar and broadly sympatric G. c. australis by a shorter head and deeper head and body, and from the morphologically similar and sympatric G. omarorum by fewer lateral-line ventral rami and fewer pored lateral-line scales anterior to the first ventral ramus. We also review the geographic distributions of all eight species of the G. carapo and G. tigre clades in the southern Neotropics, showing that G. cuia is the most widespread species in the region. These results affirm the importance of understanding the structure of variation within and between species, both geographic and ontogenetic, in delimiting species boundaries.

Etymology: The specific epithet is derived from the species’ especially deep body and head, evoking the short, rounded cuia gourd used to drink the traditional mate popular throughout its range. The convention of honoring this practice in gymnotiform taxonomy is shared with G. chimarrao (chimarrão =the mate itself) and Brachyhypopomus bombilla (bombilla = the metal straw used for drinking mate). The common name “ bombilla ” is often used to describe gymnotiform fish throughout the southern Neotropics as well.

FIGURE 9. Habitat and live appearance of Gymnotus cuia.
A. The type locality of G. cuia, Lagoa Verde, Itapuã State Park, Viamão, Rio Grande, do Sul, Brazil. Photo: Diego Cognato. B. Close-up photo of the head of a live G. cuia. Photo: Will Crampton. C. Fullbody photo of of a live G. cuia. Photo: Will Crampton.